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Home , Anaclileia, Apolephthisa, Ditomyiidae, Eudicrana, Exechiini, Greenomyia

A.4. Families of SClAROlDEA

Geir E. E. SDLI, J. R. VOCKEROTHand Loi'c MATILE

Slender to moderately robust , 2.2-13.3 Adult. Head (Figs 3-15): with posterior sur- mm long (Figs 1-2). Thoracic and tibial bristles face usually more or less flattened, and in the ma- often strong. Coxae long; tibia usually with strong jority of species inserted below level of upper apical spurs (Fig. 26). Colour varied; body usu- margin of strongly arched thorax. Eyes usually ally dull yellow, brown or black, rarely brightly densely haired, rarely bare or with a few short marked; wings sometimes with conspicuous hairs, usually situated on lower part of head and markings. widely separated above, with eye bridge incom-

Fig. 4.1. fungorum (DeGeer). 50 Geir E. E. SOLI, J. R. VOCKEROTH and Loic MATILE

Fig. 4.2. winnertzi Mik. plete in some and complete eye- duced into a slender cylindrical proboscis several hridge in Paramanota Tuomikoski (Oriental). times as long as height of head in Rhynchopla- Three ocelli ~~suallypresent, variable in position; tyura de Meijere (Oriental), Meigen (Fig. median ocellus sometimes very small or absent (Fig. 15), and nlost (Fig. 12). Lahella 6);all ocelli absent only in Hesperodes Coquillett usually large and fleshy, pillow-like, with pseudo- (Nearctic) and in Syndocosia Speiser (Afrotropi- tracheae (Fig. 4), hut greatly reduced in several cal). Frons between ocelli and antcnilal bases , Metanepsia Edwards (Afrotropi- bare or haired, sometin~eswith very strong setae cal, Oriental), Chalastonepsia and Seguyola Ma- along anterior l~order;anteriorly commonly pro- tile (Afr~tro~ical);labella very long and slender duced into well demarcated frontal t~ihercle(Fig. in most Lygistorrhinidae, except in Seguyola. 3). A frontal furrow runs between thc median Cardo reduced in most Mycctophilidae, though ocellus and the frontal tubercle, though some- not in Rondaili (Fig. 7)and times partly or entirely reduced (Fig. 6). Meigen; stipes seeiningly always present and Antenna (Figs 12-15) usually inserted at mid- with some strong setae, not rarely medially fused dle of hend, sometimes well above middle e.g., (Fig. 4). Prementum son~etimespresent as a pair in Manota Willistoil and in some Keroplatidae, of small rounded sclcrites, hut usually only the varying in length from scarcely longer than head premental apodemes remain. The outline of thcse to several times as long as body; flagellum usu- apodemes varies considerably, hut in the com- ally cylindrical, so~netimesthickened basally and mon pattern the two apodeines are inedially tapering towards apex, ~~sunllywith 14 flagel- fused 2nd articulate with the first segment of the 1ome1-es- but wit11 additional very sinall terminal labellurn anteriorly; posteriorly produced into a flagellomere in some Keroplatidae and with pair of prolonged processes (Fig. 3). Lncinia large fewer than 14 flagelloineres in other Keroplati- or minute (Fig. 8), entirely reduced in most My- dac and with 9-13 in Meigen; flagel- ~eto~hiliiiae.l'alpus apparently prehensile in lonleres strongly corupressed or pectinate or l'ara~norganiellaTonnoir (Austrnlian), ~isuallywith 11otll in some Ditomyiidae and Keroplatidae (Fig. five segnlents although first two are usually very 13), and in one species of Chalastonepsia Suli short and not apparent in dry specimens, 11ut froin Japan (Kallweit 1998), usually clothed with soinetinles with fewer than five segments - seenl- short dense hairs, but sometimes wit11 short bris- ingly four in many Mycetophilinac due to reduc- tles anlong the hairs, or with long hairs as in tion of second, andlor fusion of the first two some species of Meigen, very long (Fig", 1 l), three in Bosc and several and hair-like in an Oriental species of Chalirs- related genera (loss of apical three palpal seg- tonepsia. ments), two to four in solme Neotropical species Face ~isuallybroad and setose, but very nar- of Johrunnsen, one short segment in row in some Kcroplatidae. Clypeus sometimes Metanepsia and Chalastonepsiir, and one very long completely fused wit11 face or nearly so (Fig. 5); filanleiltous segment in I,ygistorrhina Skuse; seg- more commonly a rounded, setose sclerite be low^ ments variahle in length and form, ~isuallyslen- face, in latter case probably representing the der, hut sometimes one or more broadened or lower part of a secondary divided clypeus, the swollen; specialised sensory structures presum- upper part of which is fused with face (Crampton ably always present on segment 3, ~isuallywith 1942; h/Iatile 1990). A well-developed ciharial modified (often slightly cluhbed) setae, taking punlp, attached to the lateral parts of clypeus, following forms - a median pit (Fig. 9), a median nearly always present. 1,abruln present or absent, or lateral surface patch (Fig. l O), a dorsal and bare or with some setae. ventral pit (Asindulu~n),or a longitudinal groove Mouthparts ~~s~iallyrn~ich shorter than half with a highly modified surface and 3 deep inter- hcight of head, but about as long as height of nal invagination (Fig. 11). head ill Asindztlum Latreille (Fig. 14), Antlemon Thorax (Figs 16-20) varying in form from Haliday and most Mircrorrhyncha Winnertz; pro- compressed and deep to depressed and low. rl-l~o- 52 Geir E. E. SDLI, J. R. VOCKEROTH and Loic MATILE racic sclerites varying considerably in size, shape, Lateral cervical sclerite usually triangular, and distinctness; thoracic structure used to deter- bare. Antepronoturn well-developed, usually se- mine relationships among genera and subfamilies tose, dorsally fused with postpronoturn, forming (Shaw 1948; Shaw and Shaw 1951), but not sat- a bare, collar-like structure above the neck, ven- isfactorily and therefore requiring more study. trally partly or entirely fused with proepisternum

PtP tent - sti

car sti

Figs 4.3-11. Heads. 3-4: fusca Bezzi: 3: frontal view, 4: lateral view; 5: Paratinia sciarina Mik, frontal view; 6: (Exechiopsis) clypeata (Lundstrorn), frontal view; 7: Mycomya nitida (Zetterstedt), posterior view. 8-11: palpi: 8: Leptomor~huswalkeri Curtis; 9: Phronia siebeckii Dziedzicki; 10: Paratinia sciarina Mik; 11: Cordyla fusca (Meigen) (abbreviations: antf: antennifer, ant S: antenna1 socket, a t p: anterior tentorial pit, car: cardo, cib p: cibarial pump, clyp: clypeus, fc: face, fr: frons, fr fur: frontal furrow, fr tub: frontal tubercle, lac: lacinia, Ibl: labellun~,Ibr: labrurn, lin: lingua, pip: palpornere, prernnt ap: prernental apodeme, pstr: pseudotrachea, p t p: posterior tentorial pit, sti: stipes, tent: tentoriurn). (episternum 1); suture between the two usually fork (furca 2) in form of two heavily sclerotized, distinct, but in a few genera poorly developed, in basally jointed funnels. An elongated triangular particular posteriorly. Proepimeron (epimcron process, ventral process 2, extends laterally from 1) less conspicuous, usually triangular. Proster- the furcasternum and articulates with the inner num usually strongly reduced; most distinct fea- coxal margin. ture being the heavily sclerotized fore coxal furca Metanotum narrow, bare, situated posterior (furca 1) appearing as a pair of stalked, rounded to mediotergite, laterally fused with metapleuron and flattencd interior protul>erances. by membranous area. Halter apparently with a Shape of scutum varies from evenly to highly fairly constant outline, but one or more setae arched. 111 some genera, like in the Neotropical Inay be situated just behind its base. A distinct Thoracotropis Freeman, and in Sceptonia Win- pleural suture divides inetapleuron into anterior nertz and Winnertz, strongly produced metepisternum (episternum 3) and posterior me- anteriorly, above the head, giving the gnat a tepimeron (epimeron 3). Metepistern~im~isually somewhat stooping image. Scut~im with two loilgitudinally divided, but in most genera the inorc or less distinct infrascutal sutures, an ante- two parts can not be distinguished. Hind-coxal rior parapsidal suture and a inedian transverse fork (furca 3) and ventral process 3 both well- suture (Fig. 16). Scutal vestiture highly variable, developed. ~isuallymade up of a mixture of discal setae and Anterior thoracic spiracle located between an- bristles, ine evenly dispersed or arranged in defi- tepronotum and anepisternum, co~~~monlybor- nite lines. The two genera Leiella Enderlein, 1910 dered by fine trichia; posterior spiracle located (IVeotropical) and Allactoneura de Meijere are below laterotergite and ahove nletapleuron (Fig. 16). probably ~iniquein having the discal setae dis- Wing (Figs 31-96) considerably reduced in tinctly flattened. Mediotergite usually bare, Illore size in female of one species of rarely with medially andior laterally arranged se- Meigen (from Crozet Island), in Baeopterogynn tac. Latcrotergite prominent, ovate, bare or se- Vockeroth, in some species of Keroplatinae from tose, varying from being evenly arched to Nepal, in some specimens of hot11 sexes of one stroilgly protruding, sometimes with a pro- (Nearctic) species of Boletinn Stacger and in the nounced longitudinal keel. Anepistern~imalways Afrotropical Moriniola Matile. Species of Allac- well-developed, triangular, quadrate to subquad- toneura arc peculiar in having the wings foldcd rate, in some genera greatly developed, usually in LI vespid-like manner (Fig. 59). Veins often with at the expense of the preepisternum 2. Both an- setae; n~embraneusually densely clothed with terior and posterior hasalare usually recognis- microtrichia on both sides, and often also with able; thc former not rarely incompletely sepa- few to many macrotrichia or true setae; if micro- rated from anepistcrnum by an anepisternal trichia absent, setae present on at least most of cleft; the latter sometimes fairly large, ovate and membrane. Microtrichia arranged in fine, regular heavily sclerotized. I'reepisternum 2 separated lines in . Wing venation rather from anepisternum hy anapleural suture, possi- variable and much used in classification. Vena- bly amalgamated with the mesostern~in~ven- tion slig11tly reduced in e.g., Acnemin Winnertz trally. (This coinhined structure is frequently, but and Mik (Fig. 49), strongly rcduced erroneously, referred to as the karepisternum). in Lygistorrhinn (Fig. 4 1 ), Manotn Williston (Fig. Sllapc of preepisternum 2 fairly constant, usually 42), and Azana Walkcr (Fig. 47). Iargcr than or about as large as thc ancpistcrnuin, Heavily sclerotized portion of costa, C, ex- usually bare, but may have small setae on its tends from wing base to apex of Ri, or somc- Iowcr part. Posterior portion of mesopleuron, wllere hctwcen apices of Ri and M], always n~escpin~cron(cpimeron 2), usually bare, and clothed by sctac. Suhcosta, Sc, originating in thc strongly narrowed ventrally; distinction between basal portion of the wing, always traceable and an Lipper anepimeron and a lower katepimeron extends at most to the middle of thc wing, often obliterated by a secondary fusion. Mid-coxal partly or greatly reduced, at least distally; hu- 54 Geir E. E. SIZILI, J. R. VOCKEROTH and Loic MATILE meral crossvein, H, always present, running be- rarely well developed as in Macrocera (Fig. 36) tween the base of Sc and costa; a second cross- and some other Keroplatidae. Stem of median vein, Sc-R, frequently present between base of H fork and radial sector fused for a short distance and tip of Sc. in Keroplatidae (e.g., Fig. 37). Common stem of Anterior branch of the radius, RI, usually ex- median fork (M1 and M2) shorter than its tends to tip of the wing, but sometimes ends branches; rarely proximal portion of one or both about halfway, or even closer to the base. Poste- branches atrophied, or entirely reduced. rior branch of radius, the radial sector, Rs, radi- Cubitus consists of two anterior branches, ates into four principal veins, of which R2 and CuAl and CuA2, and one posterior branch, CUP, R3 are absent in (Matile 1990); Rs is of which CuAl is likely to represent a fusion of always present, while Rq, when present, ends in M4 and CuAl (Hennig 1954,1973; Matile 1990). C (e.g., Fig. 31) or in RI, in latter case enclosing a Commonly CuAl and CuAz with common stem, radial cell (e.g., Figs 38, 46). In some genera (e.g., but both may run separate from wing base, or Meigen) presence of R4 appears rather one of them (most likely CuA1) reduced; CUP unstable, and may be present on one wing only. usually weak and fold-like, located between CuAz Anterior branch of media, arculus, usually and the anal vein, never reaching wing margin. small and vague, located between distal median Anal vein with an anterior, AI, and a posterior plate and basal portion of radius. Of the 4 prin- branch, A2, of which the former may reach wing cipal veins originating from posterior median margin; in most genera only A1 distinct, but both branch, only M1 and M2 occur as free veins, of sometimes reduced. which M2 is likely to represent a fusion of the Crossvein R-M extends between posterior principal M2 and M3 (Matile 1990). Basal por- branch of radius and basal portion of median tion of media usually weak or entirely reduced, stem; crossvein bM-Cu connecting media and

Figs 4.12-15. Heads. 12: Lygistowhina sanctaecatharinae Thompson (Nearctic); 13: Keroplatus clausus Coquillett (Nearctic); 14: Asindulum montanum Roder (Nearctic); 15: Cnoriste megawhina Osten-Sacken (Nearctic) (after Vockeroth 1981). A.4. Families of SClAROlDEA cubitus, apparently replacing basal portion of one or more claws greatly distorted; pulvilli ab- media in (see Matile 1990; S~li sent; empodium, if present, variable in size. 1997) (e.g., Fig. 43). In Mycetophilidae cross- Abdomen usually broadest at mid length, but vein bM-Cu has previously been intepreted as in many Keroplatidae broadest near apex. Ter- the basal portion of M (e.g., Vockeroth 1981). gites and sternites of 1-6, 1-7, or 1-8 in male and In literature on Sciaroidea either R-M or bM-Cu, 1-7 in female well-developed except for sternite or both, are frequently referred to as anterior 1 which is often reduced in size, V-shaped, some- transversal, ta, and basal transversal, tb, respec- times lacking hairs; sternites 2-6 or 2-7 in many tively (e.g., Matile 1990; Snli 1997; Chandler Mycetophilidae with a pair of submedian or sub- 1994). lateral weakly sclerotized lines and sometimes Legs (Figs 21-30) with coxae elongated and also with a similar median line (fold lines) so the stout; mid coxa of male of some species of My- sternites may be partially folded longitudinally. comya and all Echinopodium Freeman (Neot- Spiracles present below margins of tergites 1-7, ropical) with an anterior process (Fig. 24); ar- but sometinles apparently lacking in segment 1, rangement and strength of bristles and hairs of or segments 1 and 2. coxa variable. Fenlora usually slender, some- Male (Figs 97-108) often with sclerites of seg- times swollen or laterally compressed, with ves- ments 7 and 8 (tergites especially) short and tele- titure variable; mid femur in male of some spe- scoped into segment 6; terminalia usually sym- cies of Leptomorphzis Curtis with a subbasal spike. metrical, but sometimes asymmetrical (one unde- Tibiae usually slender, with vestiture variable, scribed Californian Tetragonezira Winnertz, and with short setae arranged irregularly or in regu- some Mycetophila spp.), usually directed caudally, lar rows, and usually with bristles varying in sometimes rotated clockwise through 90" or strength and arrangement (Figs 22-23). Fore more with segments 7 and 8 sharing in the rota- tibia with an anteroapical depressed area bearing tion; in Megophthalmidia crassicornis (Curtis) fine setae, arranged in one or more rows (Fig. even sternites 5 and 6 are reduced, so that the 27), or more irregularly; in Synapha depressed remaining part of the abdomen is situated more area extending up to five-sevenths length of tibia, or less below tergite 6. forward flexion of in Winnertz, Metanepsia and Chalas- the terminalia below the abdomen in the Neot- tonepsia depressed area minute, or seemingly ab- ropical Rhipidita Edwards (Diton~yiidae)is prob- sent. Mid tibia of male, more rarely of female, ably unique among . Segment 9 and sometimes with a specialised sensory area which associated structures, the terminalia, extremely is rounded or elongated, or a basal, oval swelling varied in form b~ltwith an apparently constant (Fig. 28); hind tibia sometimes with a pos- basic pattern: tergite 9, sternite 9, a pair of lat- teroapical comb of stiff setae or with a dorsoapi- eral gonocoxites each with an articulated gono- cal cleft. One apical spur present on fore tibia; stylus, a pair of submedian parameres, each prin- two apical spurs on each of mid and hind tihia, cipally articulated laterally with gonocoxites and one of which may be very short; tibial spurs ex- bearing aedeagus between them (structure of tremely short only in Dolichodactyla Freeman parameres and aedeagus often difficult to deter- (Neotropical); mid and hind tibiae each with mine), and an anus-hearing proctiger; a partly re- only one spur in Fenderomyia Shaw, Monocen- duced segnlent 10 sometimes present between trota Edwards and in the Neotropical Micra- tergite 9 and proctiger, free or more or less fused pemon Edwards. Tarsi usually slender, some- with the former. Tcrgite 9 and sternite 9 distinct times with modified hairs below or with some or partly or entirely fused with gonocoxites, ~1nd segments swollen below in female. Tarsal claws sonletinles bearing spines or processes; midven- rarely simple, usually with one or more teeth be- tral line of one or both sometimes n~embranous. low, in males of some genera of Kcroplatidae Gonocoxites separate or broadly or entirely fused thick, blunt, and serrated below; in males of ventromedially, in latter case probably by a fu- some Roletincz spp. and several related genera sion with sternite 9; sometimes with distinct 56 Geir E. E. SOLI, J. R. VOCKEROTH and Loic MATILE gonocoxal lobes distally (Fig. 100). Gonostylus Costa, Truplaya Edwards, Uytalpa Edwards, particularly varied, sometimes slender and taper- Macrowhyncha Winnertz) a well-sclerotized struc- ing, but more often with lobes or processes bear- ture (probably sperm pump) attached to its ante- ing a variety of hairs, spines, or striate areas (la- rior end in addition to strong anterior apodemes mellae) (Fig. 97). Aedeagus sometimes with long present. Parameres extremely varied, and usually anterior apodemes; in several Orfeliini ( accompanied by long parameral apodemes.

v proc 3 \- I

16

Figs 4.16-20. Details of thorax. 16: basalis (Meigen), lateral view. 17-18: left anepisternum: 17: tarnatzii (Dziedzicki); 18: Phronia braueri (Dziedzicki). 19-20: antepronotum and propleuron: 19: Platurocypta fumipennis (Buk- owski); 20: Epicypta aterrima (Zetterstedt) (abbreviations: a bas: anterior basalare, anepm: anepimeron, anepst: anepister- num, anep cl: anepisternal cleft, anpl sut: anapleural suture, a par S: anterior parapsidal suture, aprnt: antepronotum, a spr: anterior spiracle, ax cord: axillary cord, bst: basisternum, 1 cerv scl: lateral cervical sclerite, cx: coxa, furc, furca, hlt: halter, Itg: laterotergite, mr: meron, mtepm: metepimeron, mtepst: metepisternum, mtg: mediotergite, mtnt: metanotum, p bas: posterior basalare, p1 wg proc: pleural wing process, preepst: preepisternum, presc: prescutum, proepm: proepimeron, proepst: proepisternum, p spr: posterior spiracle, sc: scutum, sctl: scutellurn, st: sternite, tg: tergite, trn S: transverse suture, v proc: ventral process) (17-18 after Vockeroth 1981). A.4. Families of SClAROlDEA pp - - - . - ppp

Aedeagus and parameres suspended bctwecn the posterior projection in some Keroplatidae; each gonocoxites by gonocoxal upodemes. Hypoproct round to sac-like, well sclerotized in Bolitophili- taking a variable position from immediately above dae and Lygistorrhinidac, and in at least some the ~iedeagusand parameres to below the poste- species of Ruthe, some Neotropical rior end of tergite 9, and consisting of n pair of species in the Heterotricha-group, and in Kero- lateral unsegmented cerci and a ventral sclerite platidae (e.g., Urytalpa); more weakly sclerotized or hypoproct. Hypoproct commonly weak, some- in other Keroplatidae; highly membranous and times divided medially or fused with cerci; cercus not visible in macerated specimens in Ditomyii- usually weak but variable in form, very large in dae, some Keroplatidae, and all Mycetophilidae. Diton~~iidae,with one or more transverse rows Spermathecal ducts usunlly long, slender and tor- of short spinose setae in most Roletina spp. and tuous, extremely long in Neoplatyura flava (Mac- Creagdhubhia Chandler (Chandler 1999). quart), sometimes swollen over part of their Female (Figs 109-114) with both tergite and length, opening separately medially on sternite 9 sternite 8 present, usually well-separated, tergitc or fused a short distance before opening as in 8 rarely very short and medially divided, some- some species of Sciophila, Manota and Salgusaia times seemingly fused with tergite 9; sternite 8 Vockeroth. short or long, posteriorly bearing a pair of rounded Comprehensive accounts of the adult mor- or tapered lobes, semiarticulated or more com- phology are those of Matile (1990) (Keroplati- monly con~pletelyfused, representing gonoco- dae) and Sali (1997) (Mycetophilidae) from xites 8, each usually with posterior margin in- which lrlost of the above description is taken. vnginated. Gonapophyses 8 present as a pair of flattened outgrowths above gonocoxites 8, being Egg. Eggs ovate, elongate to slender and l>o;it- variable in shape and degree of sclerotization. shaped (e.g., Metanepsia). The chorionic struc- Between and above the gonapophyses 8 is thc la- tures of the eggsllells of 15 species of bia, n membraneous, fringe-like structure (Sxtller gnats have been studied by Plachter (1 981). He 1977). Tergitc 9 usually distinct and well-devel- distinguished between eggshells with a plastron oped, haired or bare, sometimes with a distinct and a stratified and con~plexchorion, and thosc median incision; posterolaterally is a pair of without a pl~lstro~land with n simple, one-lay- gonocoxites 9, distinct in some genera (e.g., Bo- ered chorion; none of the studied eggshells had litophila), but commonly weakly developed. respiratory horns nor specialised arcas of the Sternite 9 usually fused laterally witli tergite 9 chorion. and gonapophyses 9, commonly weakly sclero- tized and with membranous areas; some out- Larvae. The larvae of many genera and numer- growths usually surrounding opening of sper- ous species are still unknown. Larvae are usually mathecal duct (gonoporc). Tergite 10 short or cylindrical and slender, but extremely slender and absent, with or without setae; sternite 10 usually oligochaete-like in some Keroplatidae, and flat- well-developed, rarely absent or strongly re- tened and slug-like in Phronia Winnertz; with a duced, co~rlmonlywith a median groove, probably dark conical dorsal case formed from frass and serving as egg guide, witli or without setae; Iat- from larval excrement in some Phronia; covered ern1 border frequently fused with hypoproct. by slime droplets in Epicypta (Zaitzev 1984~); Cercus articulated with last sclerotized tergite, and occurring in n~ucoustubes or webs formed usually weakly sclerotized and two-segmented from salivary excretion in Dindocidiidae, Kero- with a larger basal and a smaller apical segment; platidne, and Sciophilinac (except Win- sometimes one-segmented. Cercus varying in form, nertz); possessing light-producing tissues in sev- being elongate, slender, curved, and tapering; eral species of Keroplatidae; peripneustic in Di- extremely elongate in females of the Ncarctic tornyiidae and Bolitop1lilidae, hemipneustic in species of Drepanocercus Vockeroth. Two sper- most other groups, propneustic in , mathecne probably always present, with a conical in Speolepta Edw~irdsand Acornoptera Vock- 58 Geir E. E. SIZILI, J. R. VOCKEROTH and Loic MATILE eroth (Mycetophilidae) (see Zaitzev 1979), and Head (Figs 116-120) important taxonomi- apneustic in Keroplatidae, although non-func- cally, strongly sclerotized, free, well-developed, tional spiracles apparently also present in these with slender tentorial arms, and a transverse ten- groups. The spiracular system may, however, torial bridge. Head usually bare; with a change from one larval instar to another. few dorsal setae in Ditomyiidae. Antenna rather

Figs 4.21-30. Details of legs. 21: Diadocidia (D.) fewuginosa (Meigen), tarsus of fore leg. 22-23: anterior view of fore tibia: 22: Mycomya vulgaris Garrett (Nearctic); 23: Paratinia recurva Johannsen (Nearctic). 24-25: lateral view of mid coxa: 24: Mycomya (Mycomya) flavicollis (Zetterstedt); 25: Stigmatomeria crassiconzis (Stannius). 26: Orfelia tristis (Lundstrom), dis- tal part of mid tibia. 27: Megalopelrna nigroclavatum (Strobl), apical depressed area, ventral view. 28: guttiuentris (Zetterstedt), mid tibia with sensory swelling, ventral view. 29-30: apex of left hind tibia, anterior view: 29: Exechiopsis nugax (Johannsen) (Nearctic); 30: attrita Johannsen (Nearctic). (21-23, 29-30 after Vockeroth 1981). A.4. Families of SClAROlDEA -. - large, usually one-segmented and forming an elon- war; he included keys to genera of Ditomyiidne gated, convex plate; elongate and three-segment- and . ed in Bolitophilidae and Ditomyiidae (Plachter 1979a). Labrum poorly sclerotized and fleshy Pupa. Probably distinguishable from that of (not in Ditomyiidae), supported by sclerotized other groups (except ) by having the frame that articulates with two moveable arms, combination of distinct visible spurs and leg each of which carries a fan-shaped premandible; sheaths side by side rather than overlapping; prernandihle particularly well-developed and dif- partly overlapping in Macrocerinae and a few ferentiated in Keroplatidae. Mandible lamelli- Sciophilinae ( Grzegorzek). Except form, toothed along inner margin, with 5-6 to in Bolitophilidae, Ditomyiidae and Sciaridae the several prostheca near inner basal angle, pros- pupae have a characteristic, strongly arched tho- theca long and pectinate to short and blunt. Ma- rax, and a sessile undivided anterior thoracic xilla consisting of an inner blade-like lobe and an spiracle. All pupae have six abdominal spiracles, outer oval lobe, blade-like lobe serrated along in- except for Mycetophilinae with five. Pupae ner margin and ending in a sclerotized h~lrthat found in the soil or in solid tubes are typical in lies dorsal to a basal plate-like cardo; ov~lllobe having spinulae covering the surface of the ah- with a distal circular membranous area that car- domen, and segmental rows of spines (Ditornyi- ries several papillae, strongly prolonged and api- idae and Bolitophilidae); pupae surrounded by a cally tapered in some genera, e.g., Phthinia and we17 or cocoon all have a very sn~oothcuticle. A Sciophila. In Ditomyiidae the maxilla is reduced study describing the pupae of 21 species, repre- and rather weakly sclerotized, while the mandi- senting most families, is given by Plachter ble is well-developed with large, hlunt teeth. Hy- (1 979b). popharynx consisting of two c~~rvedhorizontal processes that join in midventral line and two Biology and behaviour. Mycetophilidae are vertical processes that join horizontal processes. most abundant in humid areas, especially moist Labium reduced to a small sclerotized plate at woodland; during the day adults of many spe- base of hypopharynx. cies, especially Mycetophilinae, congregate in Body with three thoracic and nine abdominal moist dark places such as overhanging stream segments; the stated number of segments in the banks and cavities under tree roots. Many species various families varies hetween authors, see e.g., can he swept from undergrowth in woods. Lewis Hennig (1948) and Madwar (1937). Each tho- and Taylor (1 965) showed that three species in racic segment bare, or nearly so, except for two England were most active at dusk and less active ventral to ventrolateral groups of four ~tlinutese- at dawn; this behaviour may be true of many spe- tae (only two in Winnertz and Asiodi- cies and is perhaps the reason why many speci- tomyia S~ligusa;three in Symmcrus Walker (Di- mens, even of apparently rare genera such as tomyiidae) (see Krivosheina and Zaitzev 19 80)) Symmer~isand Strobl, are taken in Mal- marking the position of imaginal leg discs; ah- aise traps. Some species, among others those dominal segments bare except for a few setae, with elongated mouthparts, visit flowers (e.g., mostly near the spiracles in Ditomyiidae and in Olesen and Warncke 1989), and one species of one Phronia sp. Nine to eleven ventral creeping Mycetophila was found to be among the most welts between segments of thorax and abdomen important pollinators of one species each of in Bolitophila and Mycetophilidae; each welt Liliaceae and (Messler et al. 1980); with an armature of spicules and hooks. a number of genera have been observed to feed Comprehensive accounts of the larval stage on honeydew on leaves U. A. Downes, L. Papp, are those of Madwar (1937), Plachter (197921) pers. comm. Several species are common in and Matile (1990). Hennig (1948) gave a more caves, mines, and in smaller subterranean cavi- extended summary, also based mainly on Mad- ties, like burrows of voles (Hackman 1963). In Ge~rE E. SDLI, J. R VOCKEROTH and Loic MATILE the Holarctic Speolepta is a regular inhabitant of disturbed). Larvae of Mycetophila cingulum Meigen caves in all stages, but adults may occasionally be are reported to disperse from the host fungi hy found outside caves. Most species found in these skipping, leaping distances up to 20-fold their habitats take advantage of a stable temperature body length (Camazine 1986). In Mycetophili- and humidity in periods of drought or cold (e.g., dae a large nui-nber of genera have the pupa Kjaerandsen 1993, Kurina 1996). Some species hanging in a sparse web of salivary threads. The are also known to overwinter as adults under pupa is free in Ditomyiidae and in Bolitophila, bark or in hollow stems (Vaisanen 1981). enclosed in a dense cocoon in some Mycetophili- Larvae of many species live in fleshy or woody dae and in Docosia, and apparently enclosed in fungi; both polyphagy and oligophagy have heen a weak cocoon, which may be reduced to a few reported. Most groups of fungi, including My- threads, in other groups. xomycetes, have been reported to be infested by Several species of Hymenoptera (Ichneumoni- fungus gnats (e.g., Buxton 1954, 1960; Stubbs dae, Proctotrupidae, Braconidae) have been re- and Chandler 1978; Hackman and Meinander ported as parasites of fungus gnats (Bechev 1986; 1979; Yakovlev and Zaitzev 1990; Kurina 1991; Kolarov and Bechev 1995); in North America a Chandler 1993b; Yakovlev 1994). One species nematode has also been reported (Poinar 1992). of Boletina is known to feed on Hepaticae, and this may apply also to other species in the . Classification. The Sciaroidea are generally Larvae of several species have been found in dead considered a monophy!etic group. There is, how- . wood, under bark, or in nests of birds and squir- ever, great divergence in the ranking of the sub- rels (e.g., Docosia Winnertz and Meigen), groups by different authors. Edwards (192.5), and wasps; most or all of the larvae are probably who laid the basis for subsequent classification, mycetophagous. The larva of an undescribed recognized one family with 10 subfamilies, one of Nearctic species of Brunetti has which was the Sciarinae; subsequently Edwards been observed to feed on the surface of Pedicu- (1941) included the Macrocerinae in the Kero- laris leaves (Scropl~ulariaceae).Larvae of some platidae. Hennig (1 973) considered six of the nine Keroplatidae spin wehs and capture and feed on subfamilies recognised by Edwards in his later si-nall (Mansbridge 1933); some of papers as separate families: the Lygistorrhininae these larvae live mainly or entirely in caves and he included in the Keroplatidae, following Tuo- excrete small droplets of oxalic acid on their web mikoski (1 966c); the Manotinae (following Tuomi- which kill their prey (Matile 1993); three tropi- koski 1966a), and the Sciophilinae he placed in cal species belonging to the genera Platyceridion the Mycetophilidae, although Hennig (1948) Tollet, Proceroplatus Edwards and Truplaya are had treated the Scophilinae as a separate family. reported as being myrmecophagous (Chandler Slightly modified Edwards' classification is still in and Matile 1998; Aiello and Jolivet 1998). Some use (Hutson et al. 1980; Vockeroth 19 8 1), thougl~ Keroplatidae also have larvae that produce light today most authors treat Bolitophilidae, Ditomy- (e.g., Baccetti et al. 1987); the most famous being iidae, Diadocidiidae, Keroplatidae (with or with- those of the glowe worm Arachno- out the inclusion of Macrocerinae), Lygistor- campa which uses the light to attract the prey to rhinidae, Manotidae, Mycetophilidae and Sciari- their wehs. The larvae of the Tasmanian Pla- dae as separate families in the superfamily Sciar- narivora insignis Hickman (Keroplatidae) is en- oidea (e.g., Sobs and l'app 1988; Matile 1989, doparasitic in land planarians (Hickman 1965). 1990). Matile (1 990) divides Keroplatidae into For numerous other species the larva and its bi- three subfamilies Aracl~noca~~~pinae,Macroceri- ology are con~pletelyunknown. nae and Keroplatinae, of which the latter is di- l'upation usually takes place in the ground hut vided into two tribes, Keroplatini and Orfeliini. some Mycetophilidae pupate in the host fungus The two subfamilies Sciophilinae (couplets (adults may remain quiescent in the pupal co- 37-94 in the present key) and Mycetophilinae coon for some time and emerge very rapidly if (couplets 95-1 22) have commonly been divided into tribes, mainly following Edwards (1925): nominotypical . Later, the subgenus Sciophilinae into Gnoristini, Leiini, Metanepsini Gymnogonia proved to be synonymous with Syn- (Afrotropical), Myconiyini and Sciophilini; My- plasta Skuse, 1890, and was raiscd to generic rank cetopliilinae into Exechiini and Mycetophilini. (Matile 1987). As Tuomikoski (1 966b) stated An additional tribe, Allactoncurini, was suggested tlint Allodiopsis as a compound genus seemingly by Sl~a~vand Shaw (1 95 l), hut has been by most was paraphyletic, it thus appears justified to give others included in the 1-eiini (see Sc~li1996). The generic rank to all liis suhgencra. delimitation of these tribes has been, and is still, Recent contributions on the higher phylogcny 1n~1ci1discussed (e.g., Vaisanen 1986; Suli 1997); of the Sciaroidea are tllosc by Hennig (1973), some characters ~~sedin tlie delineation of the Wood and Borkent (1989), Matile (1990, 1997). tribes, like the presence of setae on thc wing Matile (1 997) supposed Cecido~nyiidacto be the ~nembrane, undoubtedly represent plesiomor- sister-group of the Sciaroidea; within thc super- phic character states and thus result in para- family the phylogeny is as follows: (Ditomyiidae pliyletic groupings. This holds in particular for ((Ker~~latidae,Diadocidiidae) (Bolitopliilidae the tribcs Gnoristini and Leiini, hut also for the (Scinridae (Mycetophilidae, Lygistorrhinidae))))). Sciopliilini as diagnosed at present. The deline- A preliminary phylogeny of the Mycctophilidne ation of the three tribes Mycomyini (Vaisiincn is presented in Soli (1 997). I984), Exccl~ii~~iand Mycctopl~ilini(Tuomik- Rohdcndort (1 974) and several later authors oski 196613), however, Appears more sound. Fur- have referrcd fossils of vnrious periods from Up- thermore, we do also qucstion the n~onophylyof per Triassic onwards to several extinct genera and tlie commonly recognised sul>fnmilies in thc My- families of Fungivoroidea (= Sciaroidea); tlie cctophiliciae, but for convenience hot11 subfa~iii- oldest fossil definitely rcferahle to thc Sciaroiden lies are referrcd to in the text. as we know it todny, are those genera and species . . Following a clnssification or~g~nnllyproposed of h/lycetopliilidac described by Blagoderov h\ Tuoniikoski (1 96613, c), several authors todny (1 995, 19-97, 1998a, 13) from 1,ower Cretr~ceous, treat the various tribes in tlie Sciophilinae at the among them oncs species of the modern genus icvcl of st~bfan~ilics(e.g., Hennig 1973; Hack- Syntemnn Winnertz. The early Tertiary Baltic ninn et al. 198s; Matilc 1989; Zaitzev 1994). amber is very rich in Sciaroidea; ~111 families rec- Furthermore, Zaitzcv ( 1994) raiscd the trihe Al- ognized here (except Ditoniyiidae), and LI numbcr lactoneurini to tlic level of subfamily, nnd Vii- of recent genera have hecn reported. More than siinen (1 984) proposed a subfamily Eudicraninac 250 species liavc been described fro111 Baltic ;i~iil>cr. for Eridicrizna Locw. Until thc phylogenetic rela- tionship aniong the various groups have bee11 Distribution. Fungus gnats occur on all con- further elucidated, such a practice can not hc rcc- tinental areas except Antarctica, from northern ommended. Greenland south to Tierra del Fuego, and on Some species arc still undoubtedly incorrcctly most oceanic islands. Al~out4,500 species (Scia- assigned to genus and, especi;~llyin tlic Exechi- ridac excl.) have been described, more than ini, tlic correct assignnient of many species of 1,450 of then1 from thc Palaearctic, but many genera recently divided is inc certain. Therefore more species await description. All families in tlic the infornlation about distribution and number Sciaroidca li~ivereprcscntatives in the Palaearc- of spccies given in tlie key for cach genus is sul~ tic; Bolitophilidae are exclusive to the Holarctic ject to correction. rcgion, wliile niost of tlie remaining fnmilics have New statuses are proposed for two genera in a world wide distribution. Of tlie 103 Palaenrctic the present work, Notolopha Tuomikoski, I966 genera included in this key, all except 31 have and Myrosia Tuomikoski, 1966. 'rhesc were orig- been recorded in tlic Nearctic (Allactoncltra, Asio- inally among tlic 4 subgencra proposed by Tuo- ditomyia, Chalastonepsia, Clastobasis Skusc, ~iiikoski (1966b) for Allo~iiopsis Tuomikoski, Creagdhubhia, Grzegorzekia Edwnrds, Hetero- 1966; the two other were Gymnogonia and tlie trichir Loew, Isolzeurotrlyia Rrunetti, Larirypta 62 Geir E. E. SDLI, J. R. VOCKEROTH and Loic MATILE

Figs 4.31-42. Wings. 31: vockerothi Munroe (Nearctic); 32: Ditomyia potomaca Fisher (Nearctic); 33: Bolito- phila cinerea Meigen; 34: Diadocidia (D.) ferrugirzosa (Meigen); 35: Robsotzomyia reducta Matile et Vockeroth (Nearctic); 36: Macrocera variola Garrett (Nearctic); 37: Macrorrhyncha coxalis (Loew) (Nearctic); 38: Platyura nigriventris (Johannsen) (Nearctic); 39: Neoplatyura flava (Macquart); 40: Orfelia genualis (Johannsen) (Nearctic); 41: sanctaecathari- nae Thompson (Nearctic); 42: Manota sp. (Nearctic) (abbreviations: A1 and A2: branches of anal vein, bM-Cu: medio-cu- bital crossvein, CuAl and CuA2: anterior branches of cubitus, CUP: posterior branch of cubitus, d m p: distal median plate, H: humeral, M1 2nd M2: branches of media, RI: anterior branch of radius, R4 and RS: posterior branches of radius, R-M: radio-medial crossvein, Rs: radial sector, Sc: subcosta, Sc-R: subcostal-radial crossvein, stem v: stem vein) (31-38, 40-42 after Vockeroth 198 1). Edwards, Monoccntrota, Myrosia, Ncoclastol)iz- Vockcrotli 1980, 1981); Euceroplattis, however, sis Ostroverkliova, l'arempheriella Matile, l'hoeni- is known also fro111 the Oriental and Australasian kiella Cliandlcr, l'roceroplatus, l'seudoymosia Tuo- regions. mikoski, 1'~~rat~rlaEdwards, Setostylus Matile, T~ii/,l(~l"~,Urytalpa, Xenoplatyura Mallocli); ex- Identification. An easily availahle key, cover- cept tor Asiodito~nyia,known fro111 Japan and ing much of tlie western l'alacarctic fauna (not Russia11 Far East, and the five cxcl~~sivelyl'a- the subfamily r\;lycctopliilinnc), is supplied hy lacarctic genera Creagdhtibhia, Myrosia, Neoclas- Hutson et al. (1 980). Important keys, also cov- tol~asis,I'hocnikiclla, and Psctido ymosia, these ering Mycctophilin;ic, tliougli somewhrit out- genera arc known also fr(11ii tlie Orie~italandlor dated, arc found in Edwards (1925), Landrock the Afrotropical regions and a few of them from (1927, 1940) and Skguy (1.940). Good keys are the Neotropical region. On the contrary, nine also fou~idin Bei-Bicnko (1969), Krivosheina et Nenrctic genera have not bee11 rccordcd in tlie al. (1 986) and Zaitzev (1 994). Of these, the lat- P~llilenrctic(Aconzoptcrella Zaitzev, Adicronettra ter two are available only in Russian, but provide Vockcroth, Aphrastomyia Colier and Lane, Clu- good drawingmf the male genitalia. 111 addition zo1~1.aEdwards, Euceroplatus Edwards, C~arret- to the above keys, paperscdealing with a large tella Vockcrotli, Hespcrodcs, Loicia Vockerotli, proportion of the Palnearstic species of particu- and 1'alaeoplat)~z~rah4eunier). With the cxccp- lar genera are referred to in tlie key. A catalogue tion of Eticero/~latus,these genera arc all rather of most Pal;ie;irctic Sciaroiden (Scinridac excl.) poor in species and have not been recorded out- described up to 198 1 is found in Soos and Pnpp side tlie Nearctic or Neotropic:~] regions (see (1988).

Key to families and genera Adults 1. M and CuAl connected beyond level of crossvein H hy a distinct ohliquc to transverse or hori- zontal crossvein bM-Cu (e.g., Fig. 31), or by a brief contact or fusion of M and CLIAI;hasal portion of M (i.e., that txis~~lto junction of M witli bM-Cu or basal to fusion of R and M) distinct, weak or absent (e.g., Fig. 38) 2

- M and Cu connected at most hnsnlly at or very near level of crossvei~iH by a long lnorc or less liorizontnl crossvein hh4-CL],which meets Cu before base of posterior fork (Fig. 44); hasal por- tion of M entirely reduced 3 4

2. R4 present and at least 113 as long as R.5; Sc distinctly sclerotized for only a short distance, continuing as a weak fold that ends free (Fig. 31); wing membrane witli macrotrichia. Post- pronotuni with one or more long fine setae DITOMYIIDAE 3

- R4 presene and at most 115 as long as Ri (Fig. 39), or absent (Fig. 35); Sc ending in C or in R or ending free; wing menibranc witli or without macrotrichio. Postpronotal setne very short or absent 6 3. Flagellomeres with only dense fine Eye with deep rounded emarginntion opposite antenna1 base; eyes above separated by less than width of ocellrir triangle. A~lepister~iumwith erect hairs near upper margin Symmcrus Walker 4

- Flagellomeres with isolated suberect setae at least 112 as long ns diameter of flagellomere. Eye witli inner margin rounded or witli shallow rounded emnrginntion; eyes above separated by ~iiuclimore than width of ocellar triangle. Anepisternuni hare 5 64 Geir E. E. SDLI, J. R. VOCKEROTH and Lofc MATILE

Figs 4.43-54. Wings. 43: balioptera (Loew) (Nearctic); 44: Mycomya wulgaris Garrett (Nearctic); 45: Phthinia tanypus Loew (Nearctic); 46: Paratinia recurva Johannsen (Nearctic); 47: Azana sp. (Nearctic); 48: vernalis (Sher- man) (Nearctic); 49: Monoclona rufilateva (Walker); 50: Polylepta guttiventris (Zetterstedt); 51: Anaclileia sp. (Nearctic); 52: sayi (Aldrich) (Nearctic); 53: Baeopterogyna nudipes (male) Vockeroth (Nearctic); 54: Sciophila nouata Jo- hannsen (Nearctic) (abbreviations as in Figs 31-42) (all after Vockeroth 1981). A 4 Families of SClAROlDEA 6 5

P - p-pP

Mediotergite with setae on posterior half Symmerus (Symmerus Walker) 4 spp.; Snigusn 1973, M~~nroe1974, Zn~tzcv1978. Med~oterg~tebare Symmerus (Psilosymmerus Munroe) 6 \pp., .ill E.~\ternI'nlaearct~c; S.II~LI\.I1973, hluntoe 1 Y74, Z,i~t~ev1978. Veins Rs and M connected by a short but distinct crossvein bM-Cu. Palpus with at least two distinct segments, the ultimate more slendcr than the penultinlate Ditomyia Winnertz 8 spp.; Zaitzev 1978, 1994.

Veins Rs and M not connected hy a crossvein, touching at n point or over a sl~ortdist~l~lce (Fig. 32). l'alp~~swith only onc visible, stout segment Asioditomyia Saig~isa 1 sp., A. japo)lica (Sasnknwa), Jnpnn, Kussinn Far Enst; Snsnkawn 1963. Crossvein bM-Cu, or point of contact of M and CUAI,far before level of base of Rs; crossvein R-M distinct (Fig. 33) 0TOll.lAE Bolitophila Meigen 7 Crossvein bM-Cu beyond level of base of Rs, or Rs and M fused for a short distance; crossvein R-M distinct or obliterated by fusion vf R and M X

1x4 end~ngin C Bolitophila (Cliopisa Enderlein) 25 spp.; Stnckelherg 19692, I'ln\\m.~n~lI 97 1. R4 ending in R (Fig. 33) Bolitophila (Bolitophila Meigen) l i spp.; Stnckelberg 19693, I'l:lssmnn~i 197.5, Matile 1976. Crossveins R-M and bM-CLI both distinct. R4 ahsent (Fig. 34) 9

Croc\ve~nR-M absent bccause of contact or part~alfus~on of R and M; R4 present KLROI'LL\TIUAI~, 11 Crossveins R-M and bM-Cu forming a strnight line, both being short and vertical. Vein Sc reach- ing costa (Fig. 34) DIAI)<)CII)IIDA1: Diadocidia Ruthe 10 Crossvein R-M short but 17M-Cu distinctly longer and oblique or horizontal. Vein Sc ending free Heterotricha-group (unassigned to family) 3 ~lndrscribedspp.: 1 sp. in H~~terotrichaI.orw (so~~tl~rrnI'.LI~O~~) and 1 sp. each in 2 ~~ntIescribrdgener:l (north- ern Europe and Jnpnn rfipectivrly) (Chnndler, in prep.). Third flagellomere nt lellst four times as long as broad. Anepisternum haired above. Segments of fore tarsus slender in female Diadocidia (Adidocidia LaStovka et Matile) 2 spp., L). valida Mik, widespread and comnion, and D. trispiizosa l'olevoi, Finland; Stcickrlberg 196911, L:lStovkn and Mntile 1972, Polevoi 1996. Third flr~gellonlereat most 3.2 times as long as broad. Anepisternnm bare abovc. Segments 2-4 of fore tarsus swollen below in female (Fig. 21) Diadocidia (Diadocidia Ruthc) 6 spp.; Stackelberg 1969b, 1,nStovkn and Mntile 1972. Antenn~lestrongly compressed. Pnlpus very short, often porrect, with three seglncnts (Fig. 13). Prosternum haired at least laterally KL-:ROI'LA.I-INAE KEROI'LATINI 12 Antennae cylindrical or only n~oder~itclycornprcssed. Palpus drooping, with five seg~nenta. Prosternunl bare 16 R4 ending in RI. Laterotergite ha~red Keroplatus Bosc 8 spp., Zn~t~ev1991, Mnt~le1986.

R4 ending in C. Laterotergite bare 66 Geir E. E. SDLI, J. R. VOCKEROTH and Loic MATILE

Figs 4.55-66. Wings. 55: Megalopelma nigroclavatum (Strobl);56: nebulosus (Walker) (Nearctic);57: Allo- cotocera pulchella (Curtis); 58: obumbrata Loew (Nearctic); 59: Allactoneura sp. (Afrotropical); 60: Coelo- phthinia curta (Johannsen) (Nearctic); 61: Coelosia tenella (Zetterstedt); 62: Drepanocercus ensifer Vockeroth (Nearctic); 63: Aglaomyia gatineau Vockeroth (Nearctic); 64: Grzegorzekia collaris (Meigen); 65: Synapha tibialis (Coquillett) (Nearctic);66: plexipus (Garrett) (Nearctic) (abbreviations as in Figs 3142) (56-58, 60-63 and 65-66 after Vockeroth 1981). A.4. Families of SClAROlDEA 67

Mediotergite with triangular membranous area at base Skuse 1 sp., H. septeiztrionalls (Okada), Japan. Mediotergite uniformly sclerotized 14 Tibial setae irregularly arranged except sometimes near apex Cerotelion Rondani 2 spp., C. liizeatuin (Fabricius), western I'alaearctic, and C. racovitzai Mntile et Burghele-Balacesco, Ron~nnia and Iran. Tibial setae in regular rows on at least apical 113 15 Hind tibial setae in regular rows only on apical 112 to 113. Scutellum with marginal hairs short, in several rows Rocetelion Matile I sp., K. hunzerale (Zetterstedt), western I'alnenrctic; Matile 1988.

Hind tibia with setae in regular rows 011 almost entire tibial length. Scutelluni with marginal hairs long, in one row Setostylus Matile 1 sp., S. abdominalis (Sasakawa et Tamu), Japan; Sasakawa and Tmlu 1961. Branches of CuA slightly convergent beyond their base, then divergent (Fig. 36); wing mem- brane with or without macrotrichia. Anepisternum with at least a few strong erect hairs on upper half MACIIOCEIIINAE 3 7 Branches of CuA regularly divergent fro111 their base (e.g., Fig. 40); wing membrane without macrotrichia. Anepisternum bare or with short hairs above KEIIOPLATINAE ORFE1,IINI 19 Cervical sclerite bordered by a membranous area; antennae not longer than head and thorax together. Hind tibia with a row of macrotrichia on distal half ROBSONOMYIINI Robsonomyia Matile et Vockeroth 1 undeqcr~bedsp., Japan; T. Sa~gu~a(pers. conlm.). Head and cervical sclerite normal; antennae at most slightly shorter than body length, often longer. Tibiae without macrotrichia MACROCEIIINI 18 Anepimeron normal, reaching metapleuron; mediotergite high, at most projecting slightly back- wards from scutellum; no setae below metathoracic spiracle. Stem of median fork short (Fig. 36). Mid and hind tibia both with two spurs Macrocera Meigen 59 spp.; Stackelberg 1969c, Vockeroth 1976, Chandler 1990, Zaitzev 1994. Anepimeron not reaching metapleuron; mediotergite low, strongly projecting backwards from scutellum; a row of strong setae below metathoracic spiracle. Stem of median fork long. Mid and hind tibia each with only one spur Fenderomyia Shaw l unde~cr~hcd~p., Japan; T. Saigusa (pers. cowwn.) Mouthparts at least as long as head (as in Fig. 14) 20 Mouthparts much shorter than head 22 AI extending distinctly to wing n~argin.Anterior thoracic spiracle without setae on posterior niargln Asindulum Latreille 2 spp., A. ~zigrtamI.atreille, western I'alaearctic, and A. theodori Matile, Israel; Matile 1975b. A1 incomplete, becoming faint well before wing margin (as in Fig. 37). Anterior thoracic spiracle with short erect black setae on posterior margin 2 1 68 Geir E. E. SDLI, J. R. VOCKEROTH and Loic MATILE

Figs 4.67-78. Wings. 67: cincta (Johannsen) (Nearctic); 68: Boletina sp.; 69: Speolepta leptogaster (Winnertz); 70: Novakia sp. (Nearctic); 71: Docosia dichroa Loew (Nearctic); 72: sylvatica (Curtis); 73: colyeri Chandler; 74: Apolephthisa sp. (Nearctic); 75: dimidiata (Meigen); 76: Greenomyia joculator (Laffoon) (Nearctic); 77: Leia opima (Loew) (Nearctic); 78: Megophthalmidia occidentalis Johannsen (Nearctic) (abbreviations as in Figs 31-42) (67-68, 70-71 and 74-78 after Vockeroth 1981). A.4. Families of SClAROlDEA

-- ~--~--- .. .p

21. long, at least 113 of total length of probosc~s;labrum distinctly pointed, usually reacli- lng beyond labellum . , - Macrorrhyncha Winnertz, part 17 spp., Mntile 1975b. C

- Labellum short, at most 116 of total length of proboscis; lahrunl often extre~llelylong, not dis- tinctly pointed and not reaching beyond labellurn Antle~nonLoew 3 spp., A. hlzlicf~ryi(Imew), A. brevit~~zr~r~t~(I.oew) 2nd A. servzllz4nz (Wnlkrl-), nll western ['nlncnrctic; Mntile 19782. 22. R4 ending in RI;base of M weak hut distinct (Fig. 38). Empodia present, large Platyura Meigen 4 spp.; Z:iitzev 1994. - R4 ending in C (as in Fig. 39); base of M absent. Empodia absent 2 3 23. Laterotergite haired 2 4

- Laterotergite hare 2 5 24. Antennae compressed; fine tibial setae regularly arranged ~llongentire length. Wing tinted, es- pecially along veins Proceroplatus Edwards I sp., P. mikado (Okndn, 19.18). n. comb. (originnlly Zelvzira), Jnpn~i,Soutli Korea; Okadn 1938. - Antennae cylindrical; fine tibial setae irregularly arranged on hasal half. Wing clear Monocentrota Edwards -5 spp.; Zaitzev 1994. 25. Branches of M and CuA setose above 26

- Branches of M and CuA hare above 3 0 26. Posterior margin of prothoracic spiracle with a row of erect black hairs 27

- Margin of spiracle without hairs 28 27. A, well developed and reaching wing margin (Fig. 39). Tibiae with fine setae arranged in regular rows on apical 113 or 114. Mouthparts short, labrum not distinctly pinted Neoplatyura Malloch S spp.; Znit~ev1994. - AI weak and not reaching wing margin. Tibiae with tr~chiairregularly arranged along entire length. Mouthparts long, lahrum sharply pointed Macrorrhyncha Winnertz, part See couplet 2 1. 28. Fine tibial setae in regular rows. Anepisternum bare Isoneuro~nyiaBrunetti 3 spp., I. ebriold I'lassrnnnn, Germnny, 1. sctilirltfd (Mcigcn), widespread, and I. vitripeizizis (Mri~en),wrhterIi Pnlaenrctic.

- Fine tibial set~ieirreg~~larly arranged escept sometinles near tibial apex. A~~episternumhare or with short suberect hairs on upper half 2 9 29. A1 reach~ngwing margin Urytalpa Edwards 7 \pp.; Z.iitzrv 1994.

- iil extending at most 314 of distance from its base to wing margin Pyratula Edwards 5 5pp. 70 Geir E. E. SOLI, J. R. VOCKEROTH and Loic MATILE

Figs 4.79-90. Wings. 79: Zmpleta consorta Plassmann; 80: Palaeodocosia janickii (Dziedzicki);81: Hadroneura rutila (Sher- man) (Nearctic); 82: Dziedzickia marginata (Dziedzicki); 83: ciliata Winnertz; 84: Exechia attrita Johannsen (Nearctic); 85: Exechiopsis nugax (Johannsen) (Nearctic); 86: Exechiopsis (X.) mernbranacea (Lundstrom); 87: triangularis Shaw (Nearctic);88: (A.) ornaticollis (Meigen);89: vulgaris (Dziedzicki);90: Phronia cordata Lundstrom (abbreviations as in Figs 3142) (81, 83-85 and 87-90 after Vockeroth 1981). A.4. Families of SClAROlDEA 71

Tibia witli about 6 rows of setae much more closely set than others, appearing as conspicuous black lines (Fig. 26); med~oterg~tewith many short hairs Orfelia Costa 18 spp.; Zaltrev 1994. All rows of tibial setae alike; mediotergite bare 3 1 Tibial setae irregular on about basal half of tibia, in even rows on remainder. Posterior margin of anterior spiracle without hairs; metepisternum bare 3 2 Tibial setae in even rows on entire length of tibia. Posterior niargin of prothoracic spiracle with or without black hairs 3 3 Costa produced well beyond tip of vein Rs; a few short black frontal setae close to bases of antennae Xenoplatyura Malloch 4 spp., Israel; Chandler 1994b. Costa ending ;it tip of vein Rj; no sucli frontal setae Truplaya Edwards 1 sp., T. fini (,handler, Israel; Chandler 1994b. Posterior margin of anterior spiracle with erect black hairs; inetepisternuni witli dense cluster of short setae on posteroventral portion. Anal vein present Rutylapa Edwards l sp., K. ruficonzis (Zetterstedt), western I'alacarctic. Posterior margin of anterior spiracle without such hairs; anal vein absent Laurypta Edwards 2 spp., L. exserta (Ostroverkhova), Russian Far East, and L. tripotini Matile, South Korea. Rs and R1 separated from level of crossvein H; stem of h41 and Mz absent; CuAl and branches of M present as detached veins on distal part of wing (Fig. 42). Mouthparts long and slender, several times as long as height of head (Fig. 12) L.YCISTOK!IHINIL>AE Lygistorrhina Skuse I sp., L. pictij~ennis Okada, Japan; Okada 19.17. Rs arising from R well beyond crossvein H; stem of M1 and M1 present or absent. Mouth-parts usually much shorter than head; if moutliparts long and slender, both M and CuA entire and iiornially forked 3 5 Eyes meeting in a narrow eye bridge abovc antennae (except in in which eyes are reduced in hoth sexes and wings and halteres are lacking in female). Wings with stem and fork of M subequal in length, and with fork distinctly bell-shaped. Abdomen broadly inserted on thorax, covering a large proportion of the mediotergite SCIAKIDAF. Chapter 2.6 Eyes not meeting above antennae. Wings ~~suallywith fork of M much longer than stem, and lanceolate rather than bell-shaped. Abdonien narrowly inserted on thorax, most of niediotergite left free MYCETOPH1LIL)AE 36 Wing membrane either with niicrotricliia irregularly arranged (Fig. 95) and with macrotrichia present or absent, or witli microtrichia absent and niacrotrichia abundant; Sc variable, ending in C or in R or ending free; R4 present or absent. Laterotergite haired or bare. Ocelli variable in position, often far from eye margins 3 7 Microtrichia always present and, espec~allynear wing niargin, arranged in more or less regular longitudinal lines (Fig. 96); niacrotrichia usually absent, at most a few present in anal area; Sc ending free or In R; R4 absent. Laterotergite haired. Lateral ocelli touching eye margins 95 72 Geir E. E. SIZILI, J. R. VOCKEROTH and Loic MATILE

37. Fine tibial setae arranged in regular longitudinal rows (Fig. 22). Wing membrane without macro- trichia 3 8 - Fine tibial setae irregularly arranged (Fig. 23). Wing membrane with or without macrotrichia 48 38. Stem of M absent; branches of M present as detached veins on distal part of wing; R4 missing (Fig. 42). Head with a row of strong, straight, posteriorly directed bristles behind eye; three ocelli Manota Williston 1 sp., M. unifurcata Lundstrom, widespread but rare. - Stem of M present, branches of M complete; R4 present or absent. Head without strong pos- teriorly directed bristles behind eye; two ocelli 3 9 39. Vein R4 absent; C produced well beyond tip of R5 Parempheriella Matile 1 sp., South Korea (Matile, in prep.). - Vein R4 present; C ending in, or produced a short distance beyond tip of Rs 4 0 40. C extending slightly beyond apex of Rs; Rs reaching wing margin slightly before wing apex; wing membrane with a false vein between Rs and Mi; wing with conspicous dark markings (Fig. 43) Neoempheria Osten-Sacken 18 spp.; Vaisanen 1982. - C ending at apex of Rs; Rs reaching wing margin at wing apex; wing membrane without false vein between R5 and M1; wing unmarked or with obscure clouding (Fig. 44) Mycomya Rondani 41

95 96 Figs 4.91-96. Wings. 91: Mycetophila unipunctata Meigen; 92: Epicypta scatophora (Perris); 93: ornata Loew (Nearctic); 94: Sceptonia johannseni Garrett (Nearctic). 95-96: anteroapical portion of wing: 95: Synapha tibialis (Coquil- lett) (Nearctic); 96: Exechia attvita Johannsen (Nearctic) (abbreviations as in Figs 31-42) (all after Vockeroth 1981). Applies to males only: Mid coxa with spur (Fig. 24) 4 2 Applies to males only: Mid coxa without spur 43 Tergites pale with dark posterior margins. Tergite 8 bare; tergite V with a forked median struc- ture (Fig. 105). Ocellar pron~inence~~sually darker than posterior part of head Mycomya (Calcomycomya VBisanen) 5 spp.; V3islnen 1984. Tergites entirely pale or dark, or dark with paler posterior margin. Tergite 8 with setae poste- riorly; tergite 9 without a forked nledian structure (as in Fig. 106). Ocellar prominence not darker than posterior part of head Mycomya (Mycomya Rondani), part 109 spp.; V3isiine11 1984. Gonocoxites not widely separated ventrally, each with small, rounded gonocoxal lobes suhme- dially (Fig. 107); tergite 9 ~~suall~with 2 groups of dark cones, often with lateral sabre-like spines Mycomya (Mycomya Rondani), part See couplet 42. Gonocoxites widely or completely separated ventrally, without such submedian gonocoxal lobes, or gonocoxal lobes strongly modified; tergite 9 without cones or sabre-like spines 44 Tergite 9 without lateral appendages 4.5 Tcrgite 9 with lateral ~~ppendages(e.g., Figs 105-106) 46 Tergite 9 without a comb, with groups of strong lateral and submedian spines; sterna1 lateral appendages with strong spines Mycomya (Coheromyia Vaisiinen) 1 sp., M. (C.) brdircieri Vlisiinen, western I'nlnenrctic; V3is3nen 1984. Tergite 9 with an apicomedian comb of setae, without groups of strong spines; sterna1 lateral appendages without strong spines Mycomya (Cymomya Vais3nen) 1 sp., M. (C:.) circllnzd~taStaeger, widespread; Viiisinen 1984. Tergite 9 without comb or combs of spines Mycomya (Neomycomya Viiisanen) 1 sp., M. (N.) fin~l~ri~t'z(Meigen), Holnrctlc; Viiisiinen 1984. Tergite 9 with at least one comb of short spines apically 4 7 Tergite 9 with one wide apical coinh Mycomya (Lycomya Viiisanen) 3 spp., M. (L.) amllreizsis V$isiinen, enstern I'alne;lrctic, nnd M. (L.) pcctinifera Edwnrds, western I'nl;~enrctic; Viiisiinen 1984. Tergite 9 with two narrow sitbmedinn apical combs Mycomya (Mycomyopsis Viiisinen) I l spp.; Viiisiinen 1984. Wing membrane with many distinct macrotrichia and usually also microtrisl~ia 4 9 Wing membrane without macrotrichia or with at most a few near posterior margin, always with dense microtrichia 6 4 Legs extremely long and slender; first tarsomere of fore leg more than twice as long as fore tibia. Mediotergite with several strong setae laterally. Stem of posterior fork very long; CuA, strongly arched (Fig. 45) Phthinia Winnertz 14 spp.; Znitzev 198417. Legs normal; first tarsomere of fore leg subequal to or shorter than fore tibia. Mediotergite with or without setae. Stem of posterior fork, if present, not particularly long; CuQll straight or slightly curved (e.g., Fig. 46) .5 0 74 Geir E. E. SOLI, J. R. VOCKEROTH and Loic MATILE

50. Laterotergite and mediotergite bare. Tibiae without distinct bristles (Fig. 23) Paratinia Mik 2 spp., P. montana Plassmann, western Palaearctic (Caucasus), and P. sciarina Mik, western Palaearctic. - Laterotergite with strong erect hairs; mediotergite usually haired at least posteriorly. Tibiae with or without distinct bristles 5 1 51. M and CuA not clearly branched but a detached branch of one of them (probably CuA1) present near wing margin; Sc short, ending free (Fig. 47) Azana Walker 5 spp.; Coher 1995. - M or CuA, or both, clearly branched; Sc long, ending in C or in R 52 52. Sc ending in R1 (Fig. 48). Mediotergite bare Syntemna Winnertz 12 spp.; Hutson 1979. - Sc ending in C. Mediotergite haired; hairs usually long and erect but sometimes very short 53

la S b ap gst aed Par

cerc ~YP gst Par gc aP aed

aed

Figs 4.97-100. Male terminalia. 97: Coelosia fusca Bezzi, dorsal view, tergal parts removed. 98: Synapha vitripennis (Meigen), dorsal view, tergal parts removed. 99: Boletina triuittata (Meigen), ventral view. 100: (Walker), dorsal view, tergal parts removed (abbreviations: aed: aedeagus, cerc: cercus, ej ap: ejaculatory apodeme, gc: gonocoxite, gc ap: gonocoxal apodeme, gc Ib: gonocoxal lobe, gst: gonostylus, hyp: hypoproct, la: lamellae, par: paramere, par ap: parameral apodeme, proct: proctiger, S b ap: setae bearing appendage (of the gonostylus), st 9: sternite 9). A.4. Families of SClAROlDEA 75

CuA unbranched 5 4 CuA branched, with anterior branch sometimes obsolete basally 55 Macrotrichia of wing membrane reflexed, directed toward wing base; R4 present or absent (Fig. 49). Posteroventral part of metepisternum with fine hairs Monoclona Mik 6 spp.; Zaitzev 1983b. Macrotrichia decumbent, directed toward wing apex; R4 absent. Metepisternum bare Winnertz 15 spp.; Zaitzev 1982a, b. Base of M1 obsolete or very weak (e.g., Fig. 52) 5 6 Base of M1 entire and distinct 5 9 Crossvein Sc-R beyond base of Rs; R4 usually present (Fig. 50) Polylepta Winnertz, part 4 spp.; Bechev 1990b. Crossvein Sc-R, if present, before base of Rs; R4 absent 5 7 C at least one-third of the distance between apex of Rs and apex of M1; R5 moderately sinuate; crossvein R-M about twice as long as Rs (Fig. 51) Anaclileia Winnertz 5 spp.; Becl~ev19903; Zaitzev 1994. C produced at most one-fifth of the distance between apex of R5 and apex of Ml; Rj strongly sinuate; crossvein R-M about as long as Rs (Figs 52-53) 5 8 Tibiae with distinct setae about as long as tibial diameter. Anepisternum and metepisternum bare Neuratelia Rondani l0 spp.; Matile 1974. Tibiae without distinct setae. Upper part of anepisternum, and metepisternum, with fine hairs. Only male known; female possibly stenopterous Baeopterogyna Vockeroth I sp., U. irlihalyii Matile, Hungary; Matile 197.5~. Point of furcation of CuA beyond point of furcation of M (Figs 54-55); CuAl sometimes ob- solete basally 60 Point of furcation of CuA before point of furcation of M (Figs 50, 56-58); CuA1 entire 61 Macrotrichia decumbent, directed toward wing apex; macrotrichia abundant, each more than ten times as long as one microtrichium (Fig. 54). Anepisternum haired above Sciophila Meigen 48 spp.; Zaitzev 1982c, 1994. Macrotrichia of wing membrane erect or slightly reflexed, directed toward wing base; macro- trichia sparsely distributed, each about three times as long as one inicrotrichium (Fig. 55). Anepisternum bare Megalopelrna Enderlein l sp., M. ~zigroclavatum(Strobl), western l'alaearctic. C not produced beyond apex of Rs (Fig. 56) Leptomorphus Curtis 4 SPP. C produced well beyond apex of Rj 62 76 Geir E. E. SOLI, J. R. VOCKEROTH and Loic MATILE

62. Crossvein Sc-R well before base of Rs (Fig. 57). Anepisternum with many long hairs Mik 1 sp., A. pulchella (Curtis), widespread. - Crossvein Sc-R beyond base of Rs. Anepisternum bare or with a few short hairs near upper margin 63

gc lat aed gc lat gst

aed

Figs 4.101-108. Details of male terminalia. 101-106: tergite 9 and proctiger: 101: Allodiopsis domestica (Meigen); 102: Notolopha cristata (Staeger) n. stat.; 103: aurivernica Chandler; 104: Exechiopsis (E.) indecisa (Walker); 105: Mycomya (Cakomycomya) pulchella (Dziedzicki); 106: Mycomya (Mycomya) flavicollis (Zetterstedt). 107-108: male ter- minalia, ventral view: 107: M. (M.) flauicollis (Zetterstedt); 108: M. (C.) pulchella (Dziedzicki) (abbreviations: aed: aedeagus, gc: gonocoxite, gc lat: gonocoxite, lateral prolongation, gc Ib: gonocoxal lobe, gst: gonostylus). Ri alnuatc; R4 ~~suallypresent, demarcat~nga am~llcell, ahout as long as w~de(Fig. 50). Thrcc ocell~present; later,11 ocell~far from eye margln Polylepta W~nnertz,part See couplet 56. Rj nearly straight; 1x4 present, de~narcatinga snlall cell distinctly longer than widc (Fig. 58). Two ocelli present, touching eye rnnrgins Eudicrana L>ocw 2 5pp.. E. affiuis Okndn, j.lpcln, and E. ~zigriccps(I,undstriim), western I'al:~enrctic. Wings longit~~dinallyfolded; radial veins densely setose (Fig. 59). Hend with a row of strong, rccurved, posteriorly directed hristles hehind eyes. Scuturn and ahdomen covered hy narrow hut distinctly flattened scalc-like setae Allacto~leurade Meijcre I s17., A. I.ISS~I~~L>~ZS~SZnitzcv, RLISS~~IIFnr East; Znitzev 198 1.

Wing flat; radial veins with a single row of setae. Hcad with at most 3 few posteriorly directed bristles. Scuturn and abdomen with normal setne 6.5 Mediotergite with long erect hairs near posterior end. CuA2 widely divergent from CuAl (Fig. 60) Coclophthillia Edwards 1 sp., c:. thordcicd (Willllcrtz), \vestern P:ilaenrctic.

Med~otcrgiteb'1r-e. Cuhz only sliglltly divergent from CuAl (7 6 Laterotergite bare 67 Latcrotcrgite haired; hairs 1 ~~suallylong and ahundnnt hut sometilnes short and few in nunlher and confined to posterior declivity of laterotergite 8 3 Sc ending in C 6 8 Sc ending free or in R1 78 l'oint of furcation of CuA distinctly hcyond lcvcl of polnt of furcation of h{; crossvein Sc-l1 abscnt (Fig. 6 1 ) Coelosia Wi~l~lertz 1 I $pp.; So11 1'997. Point of furcation of CuA before, hclow or very sligl~tlyheyond point of furcation of M; cross- vein Sc-R present or :11>sent 69 Mouthparts forming a long slender proboscis that is several times as long as height of head (Fig. 1 S) Gnoristc Meigcn 7 spp.; Zaitzev 1994. h4outhparts shorter than height of head 70 Point of furcation of CuA very near wing hase; R4 present (Fig. 62) Drepa~locercusVockcroth I sp., l). spi~zistylttsSnli, Nor\vuy, Sweden; Se~li 1993. Point of furcation of CLIAwell beyond wing base; R4 present or absent. Female cercus usu3lly short, weakly sclerotized 7 1 Crossvein Sc-R present, at apical fourth of Sc Crossvein Sc-R near middle of Sc. or absent 76 78 Geir E. E. SOLI, J. R. VOCKEROTH and Loic MATILE

cerc

Figs 4.109-114. Female terminalia. 109-110: Coelosia tenella (Zetterstedt): 109: lateral view, 110: ventral view. 111-112: Phthinia winnertzi Mik: 111: lateral view, 112: ventral view. 113-114: Pavatinia sciarina Mik: 113: ventral view, sternal parts removed, 114: dorsal view, sternal parts (abbreviations: cerc: cercus, gc: gonocoxite, gnp: gonopore, gp: gonapophysis, hyp: hypoproct, lb: labia, no: notum, sp d: spermathecal duct, st: sternite, tg: tergite). A.4. Families of SClAROlDEA 79

Stem of median fork shorter than or subequal to crossvein R-M; point of furcation of CuA before base of crossvein R-M (Fig. 63) Aglaomyia Vockeroth l sp., A. iizgrica (St'lckelberg), widespread but rare; Stdckelberg 1948. Stem of median fork at least twice as long as crossvein R-M; point of furcatioil of CuA beyond base of crossvein R-M (Figs 64-65) 7 3 Base of posterior fork well beyond base of stem of median fork. Lateral ocelli less than their diameter from the eye margin. Empodia well-developed. R4 present or absent (Fig. 65) Synapha Meigeil 2 spp., S. fasciata Meigen and S. vitripetznis (Meigen), both western I'alaearctic. Base of posterior fork close to or before base of stem of median fork. Lateral ocelli at about their diameter distant from the eye margin. Empodia not developed. R4 always present 74 Sc setose above. R4 closely approximated to Rs and radial cell narrow or almost obliterated (Fig. 64). Setae present on stem of posterior fork Grzegorzekia Edwards 1 sp., C;. collaris (Meigen), western Palaenrctic. Sc bare. R4 further removed from Rs, fornling a larger radial cell. Setae present or absent on stem of posterior fork 75 Stem of posterior fork setose. Rs and R4 forming a rectangu1;ir radial cell. Male with sensory area on nlid tibia and spinose setae on cerci Creagdhubhia Chandler l sp., C. mrzllochortin~Chandler, Scotland; Ch.undler 1999. Stem of posterior fork bare. Rs and R4 forming a smaller trapezoidal radial cell. Male without sensory area on mid tibia and cerci without spines Phoenikiella Chandler 1 sp., 1'. phoenix (Vlisznen), Tunisin, Israel and Malta; Chandler 1999. R4 present (Fig. 66) Acomoptera Vockeroth 2 spp., A. difficilis (Dziedzicki), western I'alaearctic (rare) and A. sinica Wu et Yang, Mongolia; \Vu ancl Y:uig 1990. R4 absent 7 7 Sc ending before level of base of crossvein R-M (Fig. 67). Metepistel rnum with very short hairs that are dark in female but pale and ii~conspicuousin male Saigusaia Vockeroth 1 sp., S. flaviventrrs (Strobl), widespread. Sc ending beyond level of base of crossvein R-M (Fig. 68). Metepisternum bare Boletina Staeger, part 82 spp; Zaitzev 1994. Crossvein R-M oblique; RI more than six times as long as crossvein R-M (Fig. 69); R4 present. Apical spur of fore tibia, and longer spur of mid tibia, not longer than kipical tibial diameter Speolepta Edwards l sp., S. leptogaster (Winnertz), Holarctic. Crossvein R-M nearly horizontal, R, less than four times as long as crossvein R-M (Figs 70-73). Apical spur of fore tibia, and longer spur of mid tibia, at least twice as long as apical tibial diameter 7 9 Basal section of Rs indistinguishable because of crowding of radial veins towards C (Fig. 70) Novakia Strobl 2 spp., N. scatopsiformis Strobl, Balkan and Tunisia and N. sit~rillitnaStrobl, Austria; Z,~itzev1994. Basal section of Rs distinct 8 0 8 0 Geir E. E. SOLI, J. R. VOCKEROTH and Loic MATILE

80. Flagellum pectinate, at least in males. Palpus with one visible segment. Anteroapical depressed area very weakly developed Chalastonepsia S~li 1 sp., C. hokkaidensis Kallweit, Japan; Kallweit 1998. - Flagellum normal, tread-like. Palpus normal, 5-segmented. Anteroapical depressed area well- developed 8 1

81. Hind coxa with many posterolateral hairs near base, then bare almost to apex. R4 absent; point of furcation of CuA well beyond wing base (Fig. 71). Lateral ocelli very near eye margins Docosia Winnertz, part 23 spp. - Hind coxa with a row of rather long setose posterolateral hairs on at least apical three-quarters. R4 present; point of furcation of CuA very near or well beyond wing base (Figs 72-73). Lateral ocelli far from eye margins 8 2

abd spr 1 abd spr 7

1- 1- lab md ant mx car

abd spr 1 121 abd spr 8 Figs 4.115-121. Larvae. 115: Mycetophila sp., general view. 116-118: Mycetophila fisherae (Laffoon): 116: mandible, 117: n~axilla,118: head capsule, ventral view. 119-121: Symmerus coqulus Garrett: 119: head capsule, dorsal view, 120: head capsule, anterior view, 121: general view (abbreviations: abd spr: abdominal spiracle, ant: antenna, car: cardo, lab: labium, md: mandible, mx: n~axilla)(after Vockeroth 1981). Sc very short and ending free. CuA with a long stem, point of furcation well beyond level of crossvein R-M (Fig. 72) Tetragoneura Winnertz 7 spp. Sc long and e~:ding in R. CuA with a very short stem, or point of furcation at wing base (Fig. 73) Ectrepesthoneura Enderlein 12 spp; Chmdler 1980, l'lnssmann 1980. Sc ending in C or, if weak apically, with apex turned toward C 8 4 Sc straight and ending free, or ending in RI V 0 RI at least four times as long as crossvein R-M (e.g., Fig. 74) 8 5 RI at most three times as long as crossvein R-M (e.g., Fig. 78) 8 6 R4 present; crossvein Sc-R absent; Sc densely setose above (Fig. 74) Apolephthisa Grzegorzek l sp., A. subil~carra(Curtis), western Palaearctic. R4 absent; crossvein Sc-R usually present; Sc ~~suallybare above, rarely with a few setae (Fig. 6 8) Boletina Staeger, part See couplet 77. RI at least twice as long as crossvein R-M; M1 detached at hase, not longer than stem of median fork; crossvein R-M oblique (Fig. 75) Rondaniella Johannsen 2 spp., K. riin~idiata(Meigen), Holnrstis, nnci K. ;apoiiica (Matsumurn), Jap.111. RI about as long as crossvei~lR-M usually shorter; M1 not or only slightly detached at base, much longer than stenl of median fork; crossvein R-M nearly horizontal (Figs 76-77) 8 7 Lateral ocelli separated from eye ~llarginsby more than twice their own diameter 8 8 Lateral ocelli separated from eye margins hy less than their own diameter 8 9 Both M2 and CuAl continue to wing margin (Fig. 76) Greenomyia Brunetti 6 spp.; Zaitzev 1982~1. Both M2 and CuAl end slightly before wing margin Neoclastobasis Ostroverkhov~i 3 spp., N. draskovitsae Matile, Hungary, N. karnijoi (S'lsnkawn), Japan and N. srbrrica Ostrovrrkhova, western Siberia; Matile 19781>, Zaitzev 19 82J. Lateral ocelli touching eye margins. Fork of CuA slightly before level of apex of Sc Clastobasis Skuse 2 spp., C. alterrzarzs (Wirlnertz), wiciesprcnd, and C. girssakowskir Znitzev, lil~ssianFar Enst; Zaitzev 1994. Lateral ocelli not touching eye ~nnrgins.Fork of CuA beyond level of' apex of Sc (Fig. 77) Leia Meigen 28 spp.; I'lassmann 1973, Znit~rv1994. RI nt most twice 3s long as crossvein R-M; crossvein K-M nearly horizontal (Figs 71, 78) 91 R1 longer, ~~suallyat least tour times as long as crossvein R-M; crossvein R-M oblique (Figs 79-82) 32

PP:, \ * Sc long, lllectlng RI In a r~ghtangle; stem of med~anfork usually weakly sclerot~zcd,shorter than or subcqual In length to crossvein R-M (Fig. 71). Lateral ocell~very near eye rnarglns Docosia W~nnertz,part See couplct 8 1. Geir E E SDLI, J R VOCKEROTH and Loic MATILE -- - -

Sc short, meeting R1 in an acute angle; stem of median fork well-sclerotized, distinctly longer than crossvein R-M (Fig. 78). Lateral ocelli separated from eye margins by more than their own diameter Megophthalmidia Dziedzicki 4 spp. Sc ending free; point of furcation of CuA below or slightly beyond base of crossvein R-M; R4 absent. Hairs of laterotergite short and weak (Fig. 79) Implcta Plassmann l sp., I. co?zsorta l'lnssninnn, Sweden; Plassmann 1978; sce also Mntile 1983. Sc ending in RI; point of furcation of CuA well before base of crossveil1 R-M; R4 present or absent. Hairs of laterotergite long and strong 9 3 R4 absent; Sc ending at or before base of Rs (Fig. 80) Palaeodocosia Meunier .3 spp., P. alpicola (Strobl), western Palaenrctic, P. flaua (Edwnrds), England and 1'. jatzickii (Dziedzicki), western Palaearctic. R4 present; Sc ending beyond base of Rs (Figs 81-82) 9 4 C extending beyond apex of R5 for 115 distance to MI (Fig. 81) Hadroneura Lundstrom 2 spp., H. kanztshatica Stnckelberg, liussian Far Enst and H. palmetti Lundsrriim, western I'alnearctic (rare); Zaitzev 1994. C extending beyond apex of R.$ for 113 distance to Ml (Fig. 82) Dziedzickia Johannsen 3 spp., D. OiloOata Ostroverkhov;i, Siberia, 11. macrtira Shinji, Japan and D. t71argit~at~z(Dziedzicki), western Pnlaearctic. Anepisternum bare or with short fine hairs 96 Anepisternum with strong bristles at least near upper margin (Fig. 1 8) 114 C ending well beyond apex of R.5 (Fig. 83) Anatclla Winnertz 42 spp.; Chandler 1977, Znitzev 19893. C ending at apex of R5 9 7 Point of furcation of CuA beyond level of point of furcation of M (Figs 84-86) 9 8 Point of furcation of CuA before or opposite level of point of furcation of M (Figs 87-88) 101 Sc ending free; crossvein R-M at least twice as long as stem of median fork, usually longer; apical half of R5 straight, and divergent from M1 (Fig. 84). Pale abdominal markings, when present, usually situated towards bases of tergites Exechia Winnertz 65 spp.; Ostroverkhova and Stackelberg 1969, Krivosheinn et al. 1986. Sc more or less distinctly ending in RI; crossvein R-M at most twice as long as stein of median fork; apical half of RS curved or straight (Figs 85-86). Pale abdominal markings broadest along posterior margins of tergites 9 9 Scutum without discal bristles. Crossvein R-M never more than twice as long as stem of median fork. Male tergite 9 entire (Fig. 103) Pseudexechia Tuornikoski 9 spp; Chandler 1978. Scutum with discal bristles well-developed. Crossvein R-M short or at least twice as long as stem of median fork (Figs 85, 86). Male tergite 9 divided (Fig. 104) Exechiopsis Tuornikoski 100 100. Apex of hind tibia distinctly oblique in lateral view, dorsal surface with large triangular shining depression at apex (Fig. 29). Apical half of Rj distinctly downturned and convergent with MI (Fig. 85) Exechiopsis (Exechiopsis Tuomikoski) 45 spp.; Ostroverkhova and Stackelberg 1969, Krivoslieina et al. 1986. - Apex of hind tibia nearly truncate in lateral view, dorsal surface at most with small shining depression (as in Fig. 30). Apical half of Rs straight, and divergent from or suhparallel with M1 (Fig. 86) Exechiopsis (Xenexechia Tuoinikoski) 11 spp.; Ostroverkliova and Stnckelberg I96Y, Krivosheinn et al. 1986. 101. Branches of M, and usually also of CuA, setulose above, especially near apex 102 - Branches of M and of CuA without setulae above 110 102. Sc ending free (as in Figs 87, 92) 103

- Sc ending in R1 (as in Fig. 88) 104 103. Mid and hind coxa each with a vertical blackish mark near apex (Fig. 25); hind tibia with pos- terior bristles on no more than apical third. Flagellum of female antenna strongly swollen basally Stigmatomeria Tuomikoski l sp., S. crassicor~is(Stannius), western l'nlaearctic; see Vockeroth 1980. - Mid and hind coxa without dark mark near apex; hind tibia with posterior bristles on most of its length. Flagellum slender in both sexes Pseudobrachypeza Tuomikoski 2 spp., 1'. heluetica (Walker), western 1';ilncarctic; and 1'. pseudoheluetica I'lassrn:~nii, Austria; Pl;issmnnn 1984. 104. Most flagellomeres shorter than wide and anepisternum haired on upper half. Male terminalia small Winnertz 10.5

- Either flagellon~ereslonger than wide or anepisternum bare 106 10.5. Fore coxa with strong black bristles on apical 112 to 113 of outer margin. AI strong to weak, but always visible and extending almost to fork of CuA Brachypeza (~rachy~ezaWinnertz) t; spp.; Zaitzev 193 l, 19871,. - Fore coxa with strong black bristles only at extreme apex. A1 absent Brachypeza (Paracordyla Tuomikoski) l sp., Ij. (l'.) obscura Wlnnertz, western I'al,ienrctic. 106. Pale markings on the abdomen situated towards the bases of the tergites Pseudorymosia Tuomikoski 2 spp., P. fouea (Dziedzicki) and 1'. optiua (Dziedzicki), both western Palaearctic. - Pale markings of abdomeil most extensive towards apices of tergites 107 107. Two proepisternal bristles and two stronger scutellar bristles present. Scutum without discal bristles Synplasta Skuse 14 spp. - Three or four proepisternal bristles and four subequal scutellar bristles present. Scutum with or without discal bristles 108 108. Anepisternum with short fine setulae on upper part. Scutum covered with fine dark setulae Myrosia Tuomikoski, n. stat. 2 spp., M. rtzaculosa (~Meigen,western I'aluearctic and M. orientalis (Zaitzev), Russian 1:nr Enst.

- Anepisternum bare. Scutuill covered with fine pale setulae 109 84 Geir E. E. SDLI, J. R. VOCKEROTH and Loic MATILE

~~ ~~ pp-P ~ ~ ~- - - P- P - - - .. ~ --P~p-~ ------

109. Flagellomeres with short stiff macrotrichia. Scutum with short and strong discnl bristles. Tergite 9 of male with one pair of long bristles (Fig. 102) Notolopha Tuomikoski, n. stat. 1 sp., N. cristdtrl (Stneger), western Ptllnearctic.

,' ,: - Flagellomeres without distinct macrotrichia. Scuturn with discal bristles well-develbpeid or rc- duced. Tergite V of male with two pairs of long bristles, of which one pair is very strong (Fig. 101) Allodiopsis Tuomikoski 7 (?) spp.; Krivosheina et al. 1986. 110. Sc ending frec; AI strong, extending heyond point of furcation of CuA (Fig. 87). Mediotergite often with short subappressed hairs at upper end of posterior declivity Rymosia Winnertz 48 spp.; Dziedzicki, 1910, Ostroverkhova and Stnckelt)erg 1969, Chandlcr 19942. - Sc ending in RI; A1 variable in length and strength. Mediotergite bare 111 11 1. Anepisternum with short hairs (Fig. 17). Basal portion of M and crossvein R-M setulose above; A1 strong, extending beyond point of furcation of CuA Tarnania Tuomikoski 5 spp. - Anepisternum bare. Basal portion of M and crossvein R-M without sctulae; AI weak, not ex- tending to point of furcation of CuA (Fig. 88) 112 11 2. Hind tibia with one or more short finc posterior bristles on apical third. Scutam with subap-

pressed bristles on most of disc Marshall " 36 spp.; Krivosheina et 31. 1986, Zaitzev 198.5, 1988b.

- Hind tibia without posterior bristles. Scuturn either with discal bristles arranged in two sublat- : . era1 stripes and sometimes also a median stripe, or without discal bristles Allodia Winnertz 1 13 113. Discal bristles minute or abscnt on at least antcrior half of scutum. Abdomen with pale colour, when present, nlore extensive towards hind margins of tergites Allodia (Allodia Winnertz) ; . 10 spp.; Zaitzev 1983a, Krivosheina et al. 1986. - Discal bristles distinct on anterior h~llfof scutum. Abdomen with palc colour, when present,

more extensive towards fore margi~lsof tergites Allodia (Brachycampta Winnertz) ; % 2.5 spp.; Zaitzev 19843, Krivosheina et al. 1986. 114. Third segment of palpus very large and swollen, much thicker than subsequent segments (Fig. 11). Antennac short and stout, with 9-13 flagellomeres. Anepinleron with a sharply delimited black mark near anterior margin Cordyla Meigen I9 spp.; Krivosheina et al. 1986. - Third segment of palpus slender, not nluch thicker than subsequent segments. Antenna slender, with 14 flagellomeres. Anepimeron without black mark anteriorly 115

, , 1 15. Anepimeron bare . -C S,, ... . . - Anepimeron with hairs and bristles

116. Longest tibial bristles about -.three - times as long as tibial diameter. Sc ending in R 1 r Dynatosoma Winnertz 19 spp.; Zaitzev 1986, 1988.1. - Tibial bristles subequal in length to tibial diametcr. Sc ending free or in R (Figs 89-90) 11 7 A.4. Families of SClAROlDEA 85

sac* ,' 117. Point of furcation of CuA before, opposite, or very slightly beyond point of furcation of M (if beyond, as in vzrlcani, cubital fork more than half as long as median fork); Sc usually ending in R (Fig. 89) Trichonta Winnertz 63 spp.; Ostroverhhova and Stackelberg 1969, GngnC 1981, Chandler 1992. - Point of furcation of CuA well beyond point of furcation of M; Sc ending free (Fig. 90) 118

118. C extending more than halfway between apex of Rj and apex of M1 7 P,- /, Macrobrachius Dziedzicki 1 sp., M. kowarzi D~iedzicki,western Palaearct~c. : ,.,'<< - C extending at most very slightly beyond apex of R5 (Fig. 90) Phronia Winnertz 97 spp.; Ostroverkhova and Stackelberg 1969, Hnckmntl 1970, GagnC 1975, Chnndler 1992. 119. CuA forked (Figs 91-92) 120

- CuA simple (Figs 93-94) 122

120. CuAl slightly divergent from M2 but parallel with or convergent toward CuA2 (Fig. 91) -, Mycetophila Meigen 153 spp.; LnStovka 1963, 1972, LnStovka nnd Kidd 1974, Krivosheinn et al. 1986, Ch,~ndler1988, 1993n. - CuAt parallel with M2 but slightly divergent from CuA2 (Fig. 92) 121 123. l'ronotum only indistinctly separated from proepisternum, with uniform short setulae (Fig. 20). Fork of CuA very slightly before fork of M; crossvein bM-Cu without setae; C not produced ,- beyond apex of Rj (Fig. 92). Abdominal sternite 2 with a pair of long setae -

122. CuAl slightly divergent from M2 (Fig. 93). Mid tibia with one or more short to long ventral

bristles Zygomyia Winnertz , l5 spp.; Chandler 1991, Zaitzev 1989b.

- CuAl parallel with M2 (Fig. 94). Mid tibia without ventral bristles Sceptonia Winnertz 15 spp.; Chandler 199 1, Bechev 1995.

Economic importance. As the ecological role sionally, fungus gnats have also infected mush- of fungus gnats is little known, their economic room farms (Sasakawa 1992). Larvae of one spe- importance can at present hardly be assessed. cies, Leia arsona Hutson, have been recorded as However, in areas where mushrooms are exten- feeding on stored root-ginger (Hutson 1977). sively used as human food, the larvae of fungus gnats are looked upon as a pest. Several of the The importance of larvae in de- most popular species of edible mushrooms (e.g., composition of organic matter is nearly com- those of Boletus, Suillus, Leccinutn and Russula) pletely unknown (for a review, see Binns 1981). may be heavily infected by fungus gnats (Dely- Numerous species, however, are thought to feed Draskovits 1974; Hackman and Meinander 1979; on fungal mycelium penetrating dead organic Krivosheina et al. 1986; Yakovlev 1988). Occa- material such as rotting trunks and branches. If 86 Geir E. E. SDLI, J. R. VOCKEROTH and Loic MATILE so, their role in the process of decomposition cently the fungus gnat fauna has proved to be a Inay be much more important than is commonly good bioindicator of undisturbed forests (0k- recognized, e.g., by carrying putrefactive micro- land 1994, 1996). Lastly, mass occurrence of the organisms into the decaying material (Stubbs and New Zealand glowe worm Arachnocampa in cer- Chandler 1978; Irmler et al. 1996). Adults of tain caves is a pop~~lartourist attraction, and is some species are important in the of thus likely to play a significant economic role on certain flowers (Messler et al. 1980). More re- a local scale.

REFERENCES Aiello, A. and Jolivet, P. 1996. Myrniecophily Lower Cretaceous of Transbaikalia.] Paleont. in Keroplatidae (Diptera: Sciaroidea). J1 hT.Y. Zh. 1995 (1): 55-63. [in Russian] ent. Soc. 104(3/4): 226-230. Blagoderov, V. A. 1997. Fungus gnats of the Baccetti, B., Crovetti, A. and Santini, L. 1987. tribe Gnoristini (Diptera, Mycetophilidae) from Light-producing organs in Keroplatus tipu- the Lower Cretaceous of Transhaikalia. Pa- loides Bosc and K. reaumuri pentophthalmus leont. J. 3 l(6): 609-6 15. [Prrleontologicheskij Giglio-Tos (Diptera: Mycetophilidae). Int. J. Zhurnal 1997(6): 44-49] Morphol. Embryol. 16: 169-1 76. Blagoderov, V. A. 1998a. Fungus gnats (Di- Bechev, D. N. 1986. Sciophila rufa Meigen (Di- ptera, Mycetophilidne) froni the Lower Cre- ptera, Mycetophiloidea) as a host for Ortlio- taceous of Mongolia. Paleont. J. 32(6): 598-604. centrus stigmaticus Holmgren (Hymenopte- [Paleontologicheskij Zhurnal 1998(6): 53-59] ra, Ichneumonidae). Acta zool. bulg. 32: 60-6 1. Blagoderov, V. A. 1998b. Fungus gnats of the Bechev, D. 1990a. Recent Holarctic species of tribes Gnoristini and Leiini (Diptera, Myce- the genus Anaclileia Meunier (Insecta, Diptera: tophilidae) from the Early Cretaceous of Mycetopliilidae). Reichenbachia 28: 67-71. Transbaikalia. Paleont. J. 32(1), 54-59. [PLz- Bechev, D. 1990b. Review of the Holarctic spe- leontologicheskij Zhunzal 1 998 (1): 5 8-62] cies of genus Polylepta Winnertz. Ent. Abh. Buston, P. A. 1954. Britisli Diptera associated 53: 179-184. with fungi. 2. - Diptera bred from Myxomy- Bechev, D. 1995. The Palaearctic species of tlie cetes. Proc. R. ent. Soc. Lond. A 29: 10-12. genus Sceptonia Winnertz (Diptera: Myceto- Buxton, P. A. 1960. Britisli Diptera associated pliilidae). Trav. sc. Univ. Plovdiv, Animalia with fungi. 3. - Flies of all families reared froni 31: 7-23. about 150 species of fungi. Entomologist's Bei-Bienko, G. Y. (ed.). 1969 (1988). Keys to mon. Mug. 96: 61-94. the of the European part of USSR. Voi- Camazine, S. 1986. Leaping locomotion in My- ume 5, Part I. Sniithsonian Institution Librar- cetopliila cinguluni (Diptera: Mycetophili- ies and The National Science Foundation. dae): prepupation dispersal mechanism. Ann. Washington D.C. 1234 pp. [translation of ent. Soc. Am. 79: 140-145. the Russian edition] Chandler, J. P. 1977. Studies of sonle fungus Bernhardt, P. 1995. Notes on the anthecology gnats (Diptera: Mycetophilidae) including nine of curta (Orchidaceae). Cunning- additions to the Britisli list. Syst. ent. 2: 67-93. hamia 4(1): 1- 8. Chandler, J. P. 1978. Notes on tlie Holarctic Binns, E. S. 198 1. Fungus gnats (Diptera, Myce- species of Pseudexecliia Tuoniikoski (Diptera, topliilidae/Sciaridae) and the role of myco- Mycetophilidae) with description of a new Brit- pliagy in soil: a review. Revue Ecol. Biol. Sol ish species. Entomologist's Rec. 90: 44-5 1. 18: 77-90. Chandler, J. P. 1980. The European and eastern Blagoderov, V. A. 1995. [Fungus-gnats of the Nearctic fungus gnats in the genus Ectrepes- tribe Sciophilini (Diptera, Mycetopliilidae) from thoneura (MYcetophilidae).Syst. ent. 5: 27-4 1. A.4. Families of SClAROlDEA

Chandler, J. P. 1981. The European and North Diptera: Mycetophilidae: Sciophilinae). Rei- American species of Epicypta Winnertz (Di- chenbachia 31: 83-91. ptera: Mycetophilidae). Ent. sand. 1 2: 3 99-21 2. Crampton, G. C. 1942. The external morphol- Chandler, J. P. 1988. Thirteen species of Myce- ogy of the Diptera. Pages 10-165, in: Guide tophila Meigen (Diptera, Mycetophilidae) to the Insects of Connecticut. First fascicle. new to the British list. Br. J. ent. nat. hist. 1: External morphology, keys to families, Tany- 139-145. deridae, Ptyc11opteridae, , An- Chandler, J. P. 1990. Notes on Macrocera isopodidae, Ti~ulidae.State Geological and Meigen (Mycetophiloidea, Keroplatidae) in- Natural History Survey of Connecticut. Bttl- cluding M. nigropicea new to Britain. Dipter- letin 64: 1-509. ists Digest 3 : 27-3 1. Dely-Draskovits, A. 1974. Systematische und Chandler, J. P. 1991. New species and additions okologische Untersuchungen an den in Un- to the British list of the fungus gnat genera garn als Schadlige der Hutpilze auftretenden Zygomyia Winnertz and Sceptonia Winnertz Fliegen. IV. Mycetophilidae. Folia ent. hung. (Diptera, Mycetophilidae). Br. J. ent. nat. (S. N.) 27: 2941. hist. 4: 143-155. Dziedzicki, H. 1910. Zur Monographic der Chandler, J. P. 1992. A review of the British Gattung Rymosia Winn. Horae Soc. ent. ross. Phronia Winnertz and Trichonta Winnertz 77: 89-104. (Diptera, Mycetophilidae). Entomologist's mon. Edwards, F. W. 1925. British fungus-gnats (Dip- Mag. 128: 237-254. tera, Mycetophilidae). With a revised generic Chandler, J. P. 1993a. The Holarctic species of classification of the family. 1i.ans. R. ent. Soc. the (DeGeer) group Lond. 1924: 505-670. (Diptera: Mycetophilidae). Er. J. ent. nat. hist. Edwards, F. W. 1941. Notes on British fungus- 6: 5-1 1. gnats (Dipt., Mycetophilidae). Entomologist's Chandler, J. P. 1993b. New rearing records of mon. Mag. 77: 21-32; 67-82. fungus gnats (Diptera: Mycetophilidae) and Gagnk, R. J. 1975. A revision of the Nearctic allied families. Dipterists Digest 13: 29-35. species of the genus Phronia (Diptera: Myce- Chandler, J. P. l994a. Rymosia Winnertz (Di- tophilidae). Trans. Am. ent. Soc. 101 : 227-3 18. ptera, Mycetophilidae), a newly recognised Gagnk, R. J. 198 1. A monograph of Trichonta elenlent of wetland faunas, with five species with a model for the distribution of Holarctic new to Britain and a key to species. Ento- Mycetophilidae (Diptera). Tech. Bull. U. S. ~nologist'sGaz. 45: 199-220. Dep. Agric. 1638: 1-64. Chandler, J. P. 199417. The fungus gnats of Is- Hackman, W. 1963. Studies on the dipterous fauna rael (Diptera: Sciaroidea, excluding Sciari- in burrows of voles (Microtus, Clethriono- dae). Israel J. Ent. 28: 1-100. nnys) in Finland. Acta zool fenn. 102: 1-64. Chandler, P. J. 1999. Creagdhubhia mallocho- Hackman, W. 1970. New species of the genus rum gen. and sp. n. (Diptera, Mycetophilidae), Phronia Winnertz (Diptera, Mycetophilidae) a remarkable new Scottish gnat with a discus- from eastern Fennoscandia and notes on the sion of its relationships. Br. J. ent. nat. hist. synonynlies in this genus. Not. ent. 50: 12: [in press] 41-60. Chandler, P. and Mntile, L. 1989. A new species Hackman, W. and Meinander, M. 1979. Di- of Platyceridion Tollet (Diptera, Keroplati- ptera feeding as larvae on macrofungi in Fin- dae) with a larva predatory in ant infested land. Annls zool. fenn. 16: 50-83. internodes of Humboldtia laurifolia Vahl. Hackman, W., LaStovka, P., Matile, L. and Studia dipterologica 5: 163-1 73. Vgisanen, R. 1988. Family Mycetophilidae. Coher, E. 1. 1995. A contribution to a revision Vol. 3, pages 220-327, in Sobs, A. and Papp, of the genus Azana \Valker, 1856 (Insecta: L. (eds): Catalogue of the Palaearctic Diptera: Geir E. E. SDLI, J. R. VOCKEROTH and Loic MATILE

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